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The aggregating anemone (''Anthopleura elegantissima''), or clonal anemone, is the most abundant
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of
sea anemone Sea anemones are a group of predation, predatory marine invertebrates of the order (biology), order Actiniaria. Because of their colourful appearance, they are named after the ''Anemone'', a terrestrial flowering plant. Sea anemones are classifi ...
found on rocky, tide swept shores along the Pacific coast of North America. This cnidarian hosts endosymbiotic
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
called
zooxanthellae Zooxanthellae is a colloquial term for single-celled dinoflagellates that are able to live in symbiosis with diverse marine invertebrates including demosponges, corals, jellyfish, and nudibranchs. Most known zooxanthellae are in the genus ''Sy ...
that contribute substantially to
primary productivity In ecology, primary production is the synthesis of organic compounds from atmospheric or aqueous carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through c ...
in the
intertidal zone The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of habitats with various species o ...
. The aggregating anemone has become a model organism for the study of temperate cnidarian-algal symbioses.


Description

The polyps of the aggregating anemone reach up to eight cm across the oral disk with approximately 100 tentacles in three or four rows around the margins of the oral disk. Most are olive to bright green (depending on the species of algal symbionts present) with tentacles tipped in pink. Individuals that live in
microhabitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s that are deficient in
photosynthetically active radiation Photosynthetically active radiation (PAR) designates the spectral range (wave band) of solar radiation from 400 to 700 nanometers that photosynthetic organisms are able to use in the process of photosynthesis. This spectral region corresponds more o ...
(PAR), such as under docks or in caves, lack symbionts and are pale yellow to white in color.


Range

Pacific coast of North America from
Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U.S., ...
, United States to
Baja California Baja California (; 'Lower California'), officially the Free and Sovereign State of Baja California ( es, Estado Libre y Soberano de Baja California), is a state in Mexico. It is the northernmost and westernmost of the 32 federal entities of Mex ...
, Mexico. Most common in protected waters.


Life history

This species of anemone is capable of reproducing both sexually and asexually. As adults, aggregating anemones release
gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce t ...
s into the water that join to form genetically unique individuals that settle on intertidal rock. This genetically distinct individual can then proliferate through
binary fission Binary may refer to: Science and technology Mathematics * Binary number, a representation of numbers using only two digits (0 and 1) * Binary function, a function that takes two arguments * Binary operation, a mathematical operation that t ...
. Some argue that this is not true reproduction but actually a form of growth. Fission is often prompted in the autumn by a decrease in the abundance of food and follows sexual spawning in summer. Over time, a single individual can generate a large colony of genetically identical
polyps A polyp in zoology is one of two forms found in the phylum Cnidaria, the other being the medusa. Polyps are roughly cylindrical in shape and elongated at the axis of the vase-shaped body. In solitary polyps, the aboral (opposite to oral) end is ...
. Because of its ability to grow in this manner, the genetic entity of a colony is potentially immortal on an ecological time scale.


Natural history


Photosynthetic symbiosis

Aggregating anemones host
endosymbiotic An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within ...
, photosynthetic algae in their tentacles, oral disk and column of the polyps. In addition to a chlorophyte, two species of
dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
''Symbiodinium'', ''S. muscatinei'' and ''S. californium'' are known to inhabit the anemone. These algae are generally referred to as
zooxanthellae Zooxanthellae is a colloquial term for single-celled dinoflagellates that are able to live in symbiosis with diverse marine invertebrates including demosponges, corals, jellyfish, and nudibranchs. Most known zooxanthellae are in the genus ''Sy ...
(dinoflagellates) and zoochlorellae (chlorophytes) and many polyps concurrently host more than one type of alga within their tissues. The rate of occurrence of each alga is determined by temperature and light regimes of anemone habitats. Zoochlorellae are typically found at higher latitudes and in deeper intertidal habitats than zooxanthellae. Of the two species of zooxanthellae that inhabit the anemone, ''S. californium'' is restricted to southern California due to its intolerance of cold temperatures whereas ''S. muscatinei'' is tolerant of a broad range of temperature and irradiance levels. The mutualistic relationship of these organisms requires adaptations of each partner. The algal symbionts convert inorganic carbon into carbohydrates for use by each partner and release oxygen to the animal host in the process. To accommodate the algae, the anemone must provide concentrated
carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
to their intracellular photosynthetic guests as well as
photosynthetically active radiation Photosynthetically active radiation (PAR) designates the spectral range (wave band) of solar radiation from 400 to 700 nanometers that photosynthetic organisms are able to use in the process of photosynthesis. This spectral region corresponds more o ...
(PAR) to fuel the photosynthetic process. This restricts the symbiotic organism to
euphotic The photic zone, euphotic zone, epipelagic zone, or sunlight zone is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological pro ...
habitats and requires consistent exposure to high levels of
ultraviolet radiation Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiation i ...
(UVR). To protect from potentially damaging UVR, the algae provide mycosporine-like amino acids (MAAs) that act as sunscreen for themselves and the host. The anemones, in turn, produce
antioxidant Antioxidants are compounds that inhibit oxidation, a chemical reaction that can produce free radicals. This can lead to polymerization and other chain reactions. They are frequently added to industrial products, such as fuels and lubricant ...
s called superoxide dismutases to protect against reactive oxygen that causes oxidative stress.


Agonism between colonies

The aggregating anemone is agonistic toward other individuals with different genetic disposition. When one colony of genetically identical polyps encounters a different genetic colony, the two will wage territorial battles. Aggregating anemones have specialized tentacles called acrorhagi that are used solely to deter other colonies from encroaching on their space. When a polyp makes physical contact with a non-clonemate, it extends the acrorhagi to attack the competing anemone with stinging cells called
nematocytes A cnidocyte (also known as a cnidoblast or nematocyte) is an explosive cell containing one large secretory organelle called a cnidocyst (also known as a cnida () or nematocyst) that can deliver a sting to other organisms. The presence of this c ...
. Acrorhagi of the attacking anemone leave behind a 'peel' of the
ectoderm The ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm (the middle layer) and endoderm (the innermost layer). It emerges and originates from t ...
and nematocysts that causes tissue necrosis in the receiving animals. A study of two colonies on a boulder removed from the shore and brought into a laboratory revealed that hostilities between neighboring colonies follow the tides. As water rushed into the tank, warrior polyps inflated their acrorhagi, tripled their body length and began reaching into an empty swath of rock between the colonies. Occasionally, a polyp from one of the colonies would move into the spatial zone between the two colonies, acting as a scout, and would be attacked by the warrior polyps of the other clone. If the scout polyp received enough stings, it would be attacked by its clonemates upon return to its own colony. The return of an attacked scout to the clone with acrorhagial peel may serve to communicate the presence and identity of neighboring clones to the interior of the colony.Ayre DJ, Grosberg RK (2005) Behind anemone lines: factors affecting division of labour in the social cnidarian Anthopleura elegantissima. Animal Behaviour 70:97–110


Predators

The few known predators of the aggregating anemone include a species of nudibranch (''
Aeolidia papillosa ''Aeolidia papillosa'', known as the common grey sea slug, is a species of nudibranch in the family Aeolidiidae.Gofas, S''Aeolidia papillosa''.In: MolluscaBase (2017). Accessed through: World Register of Marine Species (WoRMS), 26 July 2017. Dis ...
)'',
leather star The leather star (''Dermasterias imbricata'') is a sea star in the family Asteropseidae found at depths to off the western seaboard of North America. It was first described to science by Adolph Eduard Grube in 1857. Description The leather st ...
, and mosshead sculpin.


Footnotes


External links

{{DEFAULTSORT:Aggregating Anemone Actiniidae Animals described in 1835